Hostname: page-component-78c5997874-94fs2 Total loading time: 0 Render date: 2024-11-09T07:47:58.122Z Has data issue: false hasContentIssue false
  • English
  • Français

Punctuated equilibria: the tempo and mode of evolution reconsidered

Published online by Cambridge University Press:  08 April 2016

Stephen Jay Gould  and
Niles Eldredge
Stephen Jay Gould
Affiliation:
Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138
Niles Eldredge
Affiliation:
Department of Fossil Invertebrates. American Museum of Natural History, Central Park West at 79th Street, New York, N.Y. 10024
Get access Rights & Permissions [Opens in a new window]

Abstract

We believe that punctuational change dominates the history of life: evolution is concentrated in very rapid events of speciation (geologically instantaneous, even if tolerably continuous in ecological time). Most species, during their geological history, either do not change in any appreciable way, or else they fluctuate mildly in morphology, with no apparent direction. Phyletic gradualism is very rare and too slow, in any case, to produce the major events of evolution. Evolutionary trends are not the product of slow, directional transformation within lineages; they represent the differential success of certain species within a clade—speciation may be random with respect to the direction of a trend (Wright's rule).

As an a priori bias, phyletic gradualism has precluded any fair assessment of evolutionary tempos and modes. It could not be refuted by empirical catalogues constructed in its light because it excluded contrary information as the artificial result of an imperfect fossil record. With the model of punctuated equilibria, an unbiased distribution of evolutionary tempos can be established by treating stasis as data and by recording the pattern of change for all species in an assemblage. This distribution of tempos can lead to strong inferences about modes. If, as we predict, the punctuational tempo is prevalent, then speciation—not phyletic evolution—must be the dominant mode of evolution.

We argue that virtually none of the examples brought forward to refute our model can stand as support for phyletic gradualism; many are so weak and ambiguous that they only reflect the persistent bias for gradualism still deeply embedded in paleontological thought. Of the few stronger cases, we concentrate on Gingerich's data for Hyopsodus and argue that it provides an excellent example of species selection under our model. We then review the data of several studies that have supported our model since we published it five years ago. The record of human evolution seems to provide a particularly good example: no gradualism has been detected within any hominid taxon, and many are long-ranging; the trend to larger brains arises from differential success of essentially static taxa. The data of molecular genetics support our assumption that large genetic changes often accompany the process of speciation.

Phyletic gradualism was an a priori assertion from the start—it was never “seen” in the rocks; it expressed the cultural and political biases of 19th century liberalism. Huxley advised Darwin to eschew it as an “unnecessary difficulty.” We think that it has now become an empirical fallacy. A punctuational view of change may have wide validity at all levels of evolutionary processes. At the very least, it deserves consideration as an alternate way of interpreting the history of life.

Type
Research Article
Information
Paleobiology , Volume 3 , Issue 2 , Spring 1977 , pp. 115 - 151
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Literature Cited

Ager, D. V. 1973. The Nature of the Stratigraphical Record. 114 pp. John Wiley; New York.Google Scholar
Ager, D. V. 1976. The nature of the fossil record. Proc. Geol. Assoc. 87:131–159.CrossRefGoogle Scholar
Anonymous. Fundamentals of Marxism-Leninism Manual. 891 pp. Foreign Languages Publishing House; Moscow.Google Scholar
Avise, J. C. 1976. Genetic differentiation during speciation. pp. 106–122. In: Ayala, F. J., ed. Molecular Evolution. Sinauer Associates; Sunderland, Mass.Google Scholar
Avise, J. C. and Ayala, F. J. 1976. Genetic differentiation in speciose versus depauperate phylads: evidence from the California minnows. Evolution. 30:46–58.Google Scholar
Ayala, F. J. 1976. Molecular genetics and evolution. pp. 1–20. In: Ayala, F. J., ed. Molecular Evolution. Sinauer Associates; Sunderland, Mass.Google Scholar
Ayala, F. J., Tracey, M. L., Hedgecock, D., and Richmond, R. C. 1974. Genetic differentiation during the speciation process in Drosophila. Evolution. 28:576–592.CrossRefGoogle ScholarPubMed
Bettenstaedt, F. 1962. Evolutionsvorgänge bei fossilen Foraminiferen. Mitt. Geol. Staatsinst. Hamburg. 31:385–460.Google Scholar
Bush, G. L. 1969. Host race formation and speciation in frugivorous flies of the genus Rhagoletis (Diptera: Tephritidae). Evolution. 23:237–251.CrossRefGoogle Scholar
Bush, G. L., Case, S. M., Wilson, A. C., and Patton, J. L.In press. Rapid speciation and chromosomal evolution in mammals. Proc. Natl. Acad. Sci.Google Scholar
Carson, H. L. 1975. The genetics of speciation at the diploid level. Am. Nat. 109:83–92.CrossRefGoogle Scholar
Darlington, P. J. Jr. 1976. Rates, patterns, and effectiveness of evolution in multi-level situations. Proc. Natl. Acad. Sci. 73:1360–1364.CrossRefGoogle ScholarPubMed
Diamond, J. 1975. Assembly of species communities. pp. 342–444. In: Cody, M. L. and Diamond, J. M., eds. Ecology and Evolution of Communities. Harvard Univ. Press; Cambridge, Mass.Google Scholar
Dodson, M. M. 1975. Quantum evolution and the fold catastrophe. Evol. Theory. 1:107–118.Google Scholar
Dodson, M. M. 1976. Darwin's law of natural selection and Thom's theory of catastrophes. Math. Biosci. 28:243–274.Google Scholar
Drooger, C. W. 1963. Evolutionary trends in the Miogypsinidae. pp. 315–349. In: von Koenigswald, G. H. R., Emeis, J. D., Buning, W. L., and Wagner, C. W., eds. Evolutionary Trends in Foraminifera. Elsevier; Amsterdam.Google Scholar
Durham, J. W. 1971. The fossil record and the origin of the Deuterostomata. North Am. Paleontol. Conv., Chicago, 1969, Proc., H:1104–1132.Google Scholar
Eldredge, N. 1971. The allopatric model and phylogeny in Paleozoic invertebrates. Evolution. 25:156–167.CrossRefGoogle ScholarPubMed
Eldredge, N. 1974. Stability, diversity, and speciation in Paleozoic epeiric seas. J. Paleontol. 48:540–548.Google Scholar
Eldredge, N. 1974. Testing evolutionary hypotheses in paleontology: a comment on Makurath and Anderson (1973). Evolution. 28:478–481.CrossRefGoogle ScholarPubMed
Eldredge, N. 1975. Character displacement in evolutionary time. Am. Zool. 14:1083–97.Google Scholar
Eldredge, N. and Gould, S. J. 1972. Punctuated equilibria: an alternative to phyletic gradualism. pp. 82–115. In: Schopf, T. J. M., ed. Models in Paleobiology. Freeman, Cooper and Co.; San Francisco, Calif.Google Scholar
Eldredge, N. and Gould, S. J. 1974. Reply to Hecht (1974). Evol. Biol. 7:303–308.Google Scholar
Eldredge, N. and Gould, S. J.In press. Evolutionary models and biostratigraphic strategies. pp. 24–40. In: Kauffman, E. G. and Hazel, J., eds. Concepts and Methods of Biostratigraphy. Dowden, Hutchinson and Ross; Stroudsburg, Pa.Google Scholar
Eldredge, N. and Tattersall, I. 1975. Evolutionary models, phylogenetic reconstruction, and another look at hominid phylogeny. In: Szalay, F. S., ed. Approaches to Primate Paleobiology. Contrib. Primatol. 5:218–242.Google Scholar
Ferris, S. D. and Whitt, G. S. 1975. Gene duplication and functional diploidization in the Catastomidae. Genetics. 80(suppl.):30.Google Scholar
Fortey, R. A. 1974. The Ordovician trilobites of Spitsbergen I. Olenidae. Skrift. Norsk Polarinst. No. 160. 129 pp.Google Scholar
Frazzetta, T. H. 1975. Complex Adaptations in Evolving Populations. 267 pp. Sinauer Associates; Sunderland, Mass.Google Scholar
Gingerich, P. D. 1974. Stratigraphic record of early Eocene Hyopsodus and the geometry of mammalian phylogeny. Nature. 248:107–109.CrossRefGoogle Scholar
Gingerich, P. D. 1976. Paleontology and phylogeny: patterns of evolution at the species level in early Tertiary mammals. Am. J. Sci. 276:1–28.CrossRefGoogle Scholar
Gingerich, P. D.In press. d. In: Hallam, A., ed. Patterns of Evolution. Elsevier; Amsterdam.Google Scholar
Gould, S. J. 1972. Allometric fallacies and the evolution of Gryphaea. Evol. Biol. 6:91–119.Google Scholar
Gould, S. J. 1976. Ladders, bushes, and human evolution. Nat. Hist. 85(No. 4):24–31.Google Scholar
Gould, S. J. 1977. Ontogeny and Phylogeny. Harvard Univ. Press; Cambridge, Mass. In Press.Google Scholar
Gould, S. J. 1977. Eternal metaphors of paleontology. In: Hallam, A., ed. Patterns of Evolution. Elsevier; Amsterdam. In Press.Google Scholar
Hanson, N. R. 1969. Perception and Discovery. 435 pp. Freeman, Cooper and Co.; San Francisco, Calif.Google Scholar
Hallam, A. 1968. Morphology, palaeoecology, and evolution of the genus Gryphaea in the British Lias. Phil. Trans. R. Soc. London B. 254:91–128.Google Scholar
Harper, C. W. Jr. 1975. Origin of species in geologic time: alternatives to the Eldredge-Gould model. Science. 190:47–48.CrossRefGoogle Scholar
Hayami, I. 1973. Discontinuous variation in an evolutionary species, Cryptopecten vesiculosus, from Japan. J. Paleontol. 47:401–420.Google Scholar
Hayami, I. and Ozawa, T. 1975. Evolutionary models of lineage-zones. Lethaia. 8:1–14.CrossRefGoogle Scholar
Hecht, M. K. 1974. Morphological transformation, the fossil record, and the mechanisms of evolution. Evol. Biol. 7:295–303.Google Scholar
Henry, J. L. and Clarkson, E. N. K. 1975. Enrollment and coaptations in some species of the Ordovician trilobite genus Placoparia. Fossils and Strata. 4:87–95.CrossRefGoogle Scholar
Howells, W. W.In press. Homo erectus in human descent: ideas and problems. Can. Assoc. Phys. Anthrop. Symp.Google Scholar
Hurst, J. M. 1975. Resserella sabrinae Bassett, in the Wenlock of Wales and the Welsh Borderland. J. Paleontol. 49:316–328.Google Scholar
Huxley, J. 1958. Evolutionary processes and taxonomy, with special reference to grades. pp. 21–38. Uppsala Univ. Arsskr.Google Scholar
Irvine, W. 1959. Apes, Angels, and Victorians. 399 pp. Meridian Books; New York.Google Scholar
Jerison, H. J. 1973. The Evolution of the Brain and Intelligence. 482 pp. Academic Press; New York.Google Scholar
Johnson, J. G. 1975. Allopatric speciation in fossil brachiopods. J. Paleontol. 49:646–661.Google Scholar
Johnson, J. G. and Norris, A. W. 1972. Tecnocyrtina, a new genus of Devonian brachiopods. J. Paleontol. 46:565–572.Google Scholar
Johnson, J. G., Reso, A., and Stephens, M. 1969. Late Upper Devonian brachiopods from the West Range Limestone of Nevada. J. Paleontol. 43:1351–1368.Google Scholar
Kellogg, D. E. 1975. The role of phyletic change in the evolution of Pseudocubus vema (Radiolaria). Paleobiology. 1:359–370.Google Scholar
Kellogg, D. E. and Hays, J. D. 1975. Microevolutionary patterns in Late Cenozoic Radiolaria. Paleobiology. 1:150–160.CrossRefGoogle Scholar
King, M. C. and Wilson, A. C. 1975. Evolution at two levels in humans and chimpanzees. Science. 188:107–116.Google Scholar
Kitts, D. B. 1974. Paleontology and evolutionary theory. Evolution. 28:458–472.Google Scholar
Klapper, G. and Johnson, D. B. 1975. Sequence in conodont genus Polygnathus in Lower Devonian at Lone Mountain, Nevada. Geol. et Paleontol. 9:65–83.Google Scholar
Kuhn, T. S. 1962. The Structure of Scientific Revolutions. 172 pp. Univ. Chicago Press; Chicago, Ill.Google Scholar
Lande, R. 1976. Natural selection and random genetic drift in phenotypic evolution. Evolution. 30:314–334.CrossRefGoogle ScholarPubMed
Leakey, R. E. F. and Walker, A. 1976. Australopithecus, Homo erectus, and the single species hypothesis. Nature. 261:572–574.Google Scholar
Lewontin, R. C. 1974. The Genetic Basis of Evolutionary Change. 346 pp. Columbia Univ. Press; New York.Google Scholar
Lewontin, R. C. and Hubby, J. L. 1966. A molecular approach to the study of genic heterozygosity in natural populations. II. Amount of variation and degree of heterozygosity in natural populations of Drosophila pseudoobscura. Genetics. 54:595–609.CrossRefGoogle Scholar
Lespérance, P. J. and Bertrand, R. 1976. Population systematics of the Middle and Upper Ordovician trilobite Cryptolithus from the St. Lawrence lowlands and adjacent areas of Quebec. J. Paleontol. 50:598–613.Google Scholar
Lovejoy, A. O. 1936. The Great Chain of Being. 376 pp. Harvard Univ. Press; Cambridge, Mass.Google Scholar
Lovtrup, S. 1974. Epigenetics. 547 pp. John Wiley; New York.Google Scholar
MacGillavry, H. J. 1968. Modes of evolution mainly among marine invertebrates. Bijdragen Tot De Dierkunde. 38:69–74.Google Scholar
Makurath, J. H. 1974. Evolution of Appalachian gypidulid brachiopods: A reply to Eldredge (1974). Evolution. 28:481–483.Google Scholar
Makurath, J. H. and Anderson, E. J. 1973. Intraand interspecies variation in gypidulid brachiopods. Evolution. 27:303–310.CrossRefGoogle Scholar
Nei, M. 1975. Molecular Population Genetics and Evolution. 253 pp. American Elsevier; New York.Google Scholar
Ovcharenko, V. N. 1969. Transitional forms and species differentiation of brachiopods. Paleontol. J. 3:57–63.Google Scholar
Oxnard, C. E. 1975. Uniqueness and Diversity in Human Evolution: Morphometric Studies of Australopithecines. 133 pp. Univ. Chicago Press; Chicago, Ill.Google Scholar
Ozawa, T. 1975. Evolution of Lepidolina multiseptata (Permian foraminifer) in East Asia. Mem. Fac. Sci. Kyushu Univ., Ser. D Geol. 23:117–164.Google Scholar
Pilbeam, D. R. 1975. Middle Pleistocene hominids. pp. 809–856. In: Butzer, K. W. and Isaac, G. L., eds. After the Australopithecines. Mouton; The Hague.CrossRefGoogle Scholar
Raup, D. M.In press. Probabilistic models in evolutionary paleobiology. Am. Sci.Google Scholar
Rensch, B. 1960. Evolution Above the Species Level. 419 pp. Columbia Univ. Press; New York.Google Scholar
Reyment, R. A. 1975. Analysis of a generic level transition in Cretaceous ammonites. Evolution. 28:665–676.CrossRefGoogle Scholar
Robison, R. A. 1975. Species diversity among agnostoid trilobites. Fossils and Strata. 4:219–226.CrossRefGoogle Scholar
Roughgarden, J. 1973. Possibilities for paleontology. Science. 179:1225.CrossRefGoogle Scholar
Rowe, A. W. 1899. An analysis of the genus Micraster, as determined by rigid zonal collecting from the zone of Rhynchonella Cuvieri to that of Micraster cor-anguinum. Q. J. Geol. Soc. London. 55:494–547.CrossRefGoogle Scholar
Ruzhentsev, V. Y. 1964. The problem of transition in paleontology. Int. Geol. Rev. 6:2204–2213.CrossRefGoogle Scholar
Schaeffer, B. 1965. The role of experimentation in the origin of higher levels of organization. Syst. Zool. 14:318–336.CrossRefGoogle ScholarPubMed
Schopf, J. W. and Oehler, D. Z. 1976. How old are the eukaryotes. Science. 193:47–49.Google Scholar
Scott, G. H. 1974. Biometry of the foraminiferal shell. pp. 55–151. In: Hedley, R. H. and Adams, C. G., eds. Foraminifera. Academic Press; New York.Google Scholar
Scott, G. H. 1976. Foraminiferal biostratigraphy and evolutionary models. Syst. Zool. 25:78–80.Google Scholar
Simpson, G. G. 1944. Tempo and Mode in Evolution. 237 pp. Columbia Univ. Press; New York.Google Scholar
Simpson, G. G. 1953. The Major Features of Evolution. 434 pp. Columbia Univ. Press; New York.CrossRefGoogle Scholar
Simpson, G. G. 1976. The compleat palaeontologist? Annu. Rev. Earth Plan. Sci. 4:1–13.CrossRefGoogle Scholar
Sprinkle, J. 1976. Classification and phylogeny of “pelmatozoan” echinoderms. Syst. Zool. 25:83–91.Google Scholar
Stanley, S. M. 1975a. A theory of evolution above the species level. Proc. Natl. Acad. Sci. 72:646–650.CrossRefGoogle ScholarPubMed
Stanley, S. M. 1975b. Clades versus clones in evolution: why we have sex. Science. 190:382–383.CrossRefGoogle ScholarPubMed
Stanley, S. M. 1976. Stability of species in geologic time. Science. 192:267–268.Google Scholar
Sylvester-Bradley, P. C.In press. Biostratigraphical tests of evolutionary theory. pp. 41–63. In: Kauffman, E. G. and Hazel, J., eds. Concepts and Methods of Biostratigraphy. Dowden, Hutchinson and Ross; Stroudsburg, Pa.Google Scholar
Truemen, A. E. 1922. The use of Gryphaea in the correlation of the Lower Lias. Geol. Mag. 59:256–268.CrossRefGoogle Scholar
Valentine, J. W. and Campbell, C. A. 1975. Genetic regulation and the fossil record. Am. Sci. 63:673–680.Google ScholarPubMed
Van Valen, L. 1973. A new evolutionary law. Evol. Theory. 1:1–30.Google Scholar
Walker, A. 1975. Splitting times among hominoids deduced from the fossil record. Burg Wartenstein Symp. 65. Progress in Molecular Anthrop. 23 pp.Google Scholar
Wilson, A. C. 1976. Gene regulation in evolution. pp. 225–234. In: Ayala, F. J., ed. Molecular Evolution. Sinauer Associates; Sunderlan, Mass.Google Scholar
Wright, S. 1967. Comments on the preliminary working papers of Eden and Waddington. In: Moorehead, P. S. and Kaplan, M. M., eds. Mathematical Challenges to the Neo-Darwinian Theory of Evolution. Wistar Inst. Symp. 5:117–120.Google Scholar
Ziegler, A. M. 1966. The silurian brachiopod Eocoelia hemisphaerica (J. de C. Sowerby) and related species. Palaeontology. 9:523–543.Google Scholar
Zuckerkandl, E. 1968. Hemoglobins, Haeckel's “Biogenetic Law,” and molecular aspects of development. pp. 256–274. In: Rich, A. and Davidson, N., eds. Structural Chemistry and Molecular Biology. W. H. Freeman; San Francisco, Calif.Google Scholar